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(Combretaceae) from South Africa

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Phytotaxa 434 (1): 001–012 ISSN 1179-3155 (print edition) https://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2020 Magnolia Press Article ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.434.1.1 A new species of Combretum sect. Ciliatipetala (Combretaceae) from South Africa RICHARD G.C. BOON1*, MARIE JORDAAN2,3 & ABRAHAM E. VAN WYK2,3 1Biological Sciences, School of Life Sciences, University of KwaZulu-Natal, Private Bag X5400, Durban, 4000 South Africa. 2H.G.W.J. Schweickerdt Herbarium, Department of Plant and Soil Sciences, University of Pretoria, Pretoria, 0002 South Africa. 3National Herbarium, South African National Biodiversity Institute, Private Bag X101, Pretoria, 0001 South Africa. *Author for correspondence. E-mail: [email protected] Abstract Combretum eugeneanum, a new species from northeastern KwaZulu-Natal, South Africa and confined to the Maputaland Centre of Plant Endemism, is described, illustrated, mapped, and compared with southern African members of the genus with which it may be confused. In a narrowly defined genus Combretum, the new species belongs to Combretum sect. Ciliatipetala. In herbaria, it has usually been confused with close relatives C. apiculatum and C. edwardsii, as well as several other more distantly related members of the genus, in particular C. woodii. The new species is also closely related to the recently described C. stylesii. It is readily distinguished as an essentially glabrous woody climber or scrambling shrub needing other vegetation for support, leaf apices rarely apiculate, tertiary veins raised on the adaxial leaf surface, inflorescences few-flowered and subcapitate, upper hypanthium cupuliform, flowers with orange-red centres (discs) and peltate scales comprised of essentially eight radial cells, most of which are subdivided by at least one tangential wall, the resulting outer and inner cell(s) often with at least one additional radial wall. Combretum eugeneanum grows in Sand Forest and associated sandy bushveld, and its range and habitat does not overlap with that of C. edwardsii or C. stylesii, both of which are also very often lianas. Keywords: anatomy, Combretoideae, Combretum eugeneanum, endemism, Maputaland Centre of Endemism, peltate scales, scales, taxonomy Introduction Combretum Loefling (1758: 308) belongs to Combretaceae subfam. Combretoideae, tribe Combreteae (Engler & Diels 1899). The genus, when broadly defined, comprises about 276 species (POWO 2019) with a pantropical distribution mainly in tropical Africa and Asia, but absent in most of Australia and the Pacific Islands (Stace 2007; Mabberley 2017). The centre of diversity of the genus is on the African continent. If a narrow generic definition is used instead, and Quisqualis Linnaeus (1762: 556) and other segregate genera are excluded, about 252 species are included in Combretum. Members of Combretum include trees, shrubs and woody climbers. The leaves are opposite, whorled (especially ternate), sub-opposite or rarely alternate, entire, lack stipules or stipules very small; and petioles sometimes persist and form ± hooked spines. Flowers are bisexual, actinomorphic, 4- or 5-merous; the petals variously developed and coloured; the stamens twice as many as the petals, in 1 or 2 series; the ovary inferior and the fruit 4- or 5-winged (rarely an unwinged nut). In the Flora of southern Africa (FSA) region (Namibia, Botswana, South Africa, Eswatini and Lesotho), 33 species of Combretum (narrowly defined and excluding one Quisqualis species) occur (Maurin et al. 2010; Jordaan et al. 2011b). In the present contribution we follow the traditional narrow concept of the genus. Combretum consists of three subgenera (Exell & Stace 1966): Subgen. Combretum comprises species with subsessile, peltate scales, in addition to unicellular, compartmented, non-glandular hairs with a basal compartment (= combretaceous hairs), but without glandular hairs, flowers usually 4-merous (petals usually not red); Subgen. Cacoucia (Aublet 1775: 450) Exell & Stace (1966: 10) includes species without scales, but with microscopic short, capitate, stalked glands, in addition to combretaceous hairs, and flowers 5-merous, or if 4-merous then petals usually red (Exell & Stace 1966; Exell 1978); Accepted by Ronell Klopper: 5 Feb. 2020; published: 24 Feb. 2020 1 Licensed under a Creative Commons Attribution 4.0 International License http://creativecommons.org/licenses/by/4.0/ Subgen. Apetalanthum Exell & Stace (1966: 11) is monotypic and the single species has combretaceous hairs, stalked glands and scales, lacks petals, and occurs in Southeast Asia. FIGURE 1. Combretum eugeneanum. A. Trunk of a plant reaching the Sand Forest canopy at False Bay Park, Hluhluwe. B. Flowers at Tembe Elephant Park; note orange-red disc. C. Fruiting branchlet at Ndumo Game Reserve. Photograph taken in March when samaras fully grown, but before drying during autumn and early winter. Note glutinous seed pod. Photographs: R.G.C. Boon. 2 • Phytotaxa 434 (1) © 2020 Magnolia Press BOON ET AL. FIGURE 2. Combretum eugeneanum, leaf and scale morphology. A. Leaves, showing adaxial (top) and abaxial (bottom) lamina surfaces. Note the thin glutinous (shiny) secretion on the lamina surfaces and raised tertiary veins on the adaxial surface. B. Selection of scales (stained with Sudan IV) from leaves of the same collection (Ward 1970) to show variation in size, shape and number of cells. Scale bar in mm refers to leaves; one in μm to scales. Photographs: R.G.C. Boon (leaves) and A.E. van Wyk (scales). A NEW SPECIES OF COMBRETUM Phytotaxa 434 (1) © 2020 Magnolia Press • 3 FIGURE 3. Combretum eugeneanum. A. Flowering branchlet. B. Fruiting branchlet. C. Flower. D. Flower with half removed. E. Hair-tuft domatia in principal lateral vein axils of abaxial leaf blade surface. Scale bar = 10 mm (A & B), or 1 mm (C–E). A based on Ward 2644, B based on Moll 5632, C based on Ward 2661, D based on Moll 4359. Artist: Daleen Roodt. Combretum subgen. Combretum comprises 11 sections in continental Africa (Maurin et al. 2010; Jordaan et al. 2011a). The new species described here belongs to the diverse and taxonomically difficult section Ciliatipetala 4 • Phytotaxa 434 (1) © 2020 Magnolia Press BOON ET AL. Engler & Diels (1899: 32). This section is characterized by species with very small petals, 0.5–1.5 mm long, petal apices ciliate or pilose, except C. petrophilum Retief (1986: 44) and C. psidioides subsp. glabrum Exell (1968: 18), leaf apices often apiculate, flowers with disc margins short, free and pilose, fruit 4-winged, 15–30 mm long, scales relatively small, from 40–120 μm diameter, scattered, not overlapping (Maurin et al. 2010; Jordaan et al. 2011a). According to Stace (1980) this section comprises ten species and is restricted to Africa and the Arabian Peninsula. Gere et al. (2015), studying the origin of the family Combretaceae on the African continent, hypothesized that within subgenus Combretum, the most diverse section Ciliatipetala diverged ca. 13.2 mya and split into two main subclades, one comprising mainly southern African species and the other with species from the rest of Africa. Molecular phylogenetic studies done on the Combretaceae by Maurin et al. (2010) showed that there are at least three undescribed species in C. sect. Ciliatipetala. One of these, (C. sp. nov. B), from the Tugela River Valley in KwaZulu-Natal, has since been described as Combretum stylesii Maurin et al. (2011: 106). A second species from Maputaland was provisionally referred to as C. sp. nov. C in Maurin et al. (2010) and the earliest known herbarium specimen was collected by Father Jacob Gerstner in 1944 (Gerstner 4953 in PRE). Specimens of this taxon have sometimes been identified in herbaria as a possible new species of Combretum, but more frequently as C. apiculatum Sonder (1850: 45), C. edwardsii Exell (1968: 19) or C. woodii Dümmer (1913: 181), and occasionally as one of at least six other species of Combretum. The purpose of this paper is to formally describe this new species as C. eugeneanum. Materials and methods The morphological description was made by the standard herbarium procedures examining specimens of the following herbaria: BNRH, NH, NU, PCE, PRE, PRU and UDW (acronyms according to Thiers 2018). One of us (RGCB) visited the distribution area several times to make field and morphological observations. Godfrey Lang, a landowner near Hluhluwe, assisted with daily observations over eight weeks of a plant breaking dormancy and flowering in the spring of 2019. Descriptors used to indicate abundance and frequency follow Schmid (1982). The distribution map was constructed using coordinates provided on or derived from specimen labels. In the section “Additional collections (paratypes)”, locality citations were reproduced as per the specimen labels. In some cases the spelling of the locality name was corrected and is shown in square brackets. Specimens are arranged according to the Degree Reference System proposed by Edwards & Leistner (1971); the quarter degree grid reference is supplied between brackets after each locality cited. The distribution map was compiled from specimen data using ArcView 3.1 software. The original base map is based on the GTOPO30 global digital elevation model, and colours were modified in Global Mapper v6.06. A preliminary conservation assessment was conducted using the standard procedures based on IUCN guidelines (IUCN 2012). GeoCAT (Bachman et al. 2011) was used to estimate Extent of Occurrence (EOO) and Area of Occupancy (AOO) using a 2 km and 5 km cell width. To study the structure of the peltate scales, young, still conduplicate leaves were removed from herbarium specimens and rehydrated by heating in distilled water. The rehydrated but intact leaves were then stained in a 1% solution of Sudan IV in 70% ethanol for at least 24 h (Stace 1965). Following staining, the surface of the leaves was gently scraped with a razor blade to collect some scales, taking care not to cut through the surface itself. Material collected along the edge of the blade was mounted (temporarily) by suspending it in a drop of glycerol on a slide.
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